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Response of Aquatic Macroinvertebrates to a Gradient Of 1 Taxonomic and trait macroinvertebrate patterns across hydrological connectivity in 2 temperate, subtropical, Mediterranean, and arid river floodplains 3 Belinda Gallardoa,b*, Sylvain Dolédecc, Amael Paillexb,d, David B. Arscotte, Fran Sheldonf, Florencia Zillig, 4 Sylvie Mérigouxc, Emmanuel Castellad and Francisco A. Comína 5 6 a Applied and Restoration Ecology Group, Pyrenean Institute of Ecology (IPE-CSIC), Avda. Montañana 7 1005, 50192 Zaragoza (Spain) 8 b Aquatic Ecology Group, Department of Zoology, University of Cambridge, Downing St. CB2 3EJ 9 Cambridge (U.K.) 10 c Université Lyon 1; CNRS, UMR 5023, LEHNA, Biodiversité des Ecosystèmes Lotiques, F-69622, 11 Villeurbanne (France) 12 d Laboratory of Ecology and Aquatic Biology, University of Geneva, Route de Rize 7, 1207 Carouge 13 (Switzerland) 14 e Stroud Water Research Center, Avondale, PA 19311 (USA) 15 f Australian Rivers Institute, Griffith University, Nathan, Qld. 4111 (Australia) 16 g Instituto Nacional de Limnologia (UNL-CONICET), Ciudad Universitaria S/N, Ruta 168, Santa Fe 3000 17 (Argentina) 18 *Correspondence to: 19 Dr Belinda Gallardo 20 Current address: 21 Aquatic Ecology Group, Department of Zoology, University of Cambridge 22 Downing St. CB2 3EJ Cambridge (UK) 23 e-mail: [email protected] / [email protected] 24 Phone: +44 01223336617 / Fax : +44 0122336676 25 1 26 Abstract 27 Despite the wide recognition that benthic macroinvertebrates respond to changes in hydrological 28 connectivity within floodplain ecosystems, no generalizations about patterns in community structure 29 and functionality across large scales and different climates have been yet established. Using information 30 from six large rivers located in four different climate regions (subtropical, temperate, Mediterranean 31 and arid), we compared benthic macroinvertebrate responses along lateral gradients of hydrological 32 connectivity. We tested hypotheses related to differences among climate regions and to similar 33 hydrological constraints under any climate. First, multivariate ordinations demonstrated a higher 34 overlap of trait community composition (55.7% variance explained by the first two axes) than taxonomic 35 composition (19.2%) among floodplains, suggesting high inter-climate trait stability. Second, across a 36 gradient of lateral connectivity, Linear Mixed Effect (LME) models supported 7 out of 15 hypotheses, 37 which evidenced remarkably consistent macroinvertebrate patterns in floodplains regardless of climate 38 regime. Taxon and trait richness were positively related and peaked in sites with intermediate 39 hydrological connectivity. Richness of Ephemeroptera, Trichoptera, and Plecoptera peaked in highly 40 connected sites, while richness of Coleoptera, Odonata and Heteroptera was higher in the most 41 fragmented sites. Our expectations related to the feeding structure of macroinvertebrates (e.g. 42 shredders and scrapers more abundant in connected channels, and predators and deposit feeders in 43 fragmented sites) were better supported by observations than those regarding life history (e.g. 44 plurivoltinism and short life spans in connected channels). This difference was related to the influence of 45 extended periods of fragmentation (i.e. hydrological disconnection) as a complementary disturbance 46 factor to flooding in the filtering of traits. Trait similarities and dissimilarities found in this study suggest 47 that large-scale filters do operate on communities resulting in apparently different trait combinations in 48 temperate and Mediterranean floodplains when compared to arid and subtropical environments. This 49 latter finding needs to be considered to formulate ecologically meaningful a priori predictions. 50 Keywords: River Habitat Templet, Paraná River, Tagliamento River, Rhône River, Ebro River, Murray 51 River, Cooper Creek, aquatic macroinvertebrates 52 2 53 Introduction 54 Connectivity and fragmentation are fundamental concepts in ecology influencing the distribution of 55 organisms within an ecosystem (e.g. Moilanen and Nieminen 2002). Connectivity allows organisms to 56 move from an area to another, it increases the resilience of an ecosystem towards disturbance and 57 supports key ecosystem functions (e.g. Moilanen and Nieminen 2002). River floodplains comprise a 58 complex mosaic of interconnected aquatic and terrestrial patches that are arranged along a gradient of 59 lateral hydrological connectivity (e.g. Ward and Stanford 1995). This gradient of connectivity extends 60 from water bodies with a high frequency of connectivity, either due to their position adjacent to the 61 main channel (i.e. connected channels) or by being connected to the main channel at high flow levels 62 (e.g. water bodies that range in distance from the river channel), to those fragmented water bodies that 63 are only inundated under the most extreme floods (e.g. Amoros and Roux 1988). Since the first 64 definition of the Flood Pulse Concept in tropical environments (Junk et al. 1989) and its extension to 65 temperate areas (Tockner et al. 2000), there has been wide recognition that floodplain aquatic 66 communities respond to changes in hydrological connectivity in a predictable way (see the reviews of 67 Junk and Wantzen 2004, Petts and Amoros 1993). Nevertheless, to our knowledge no generalizations 68 about benthic macroinvertebrate structure and functionality across large scales and different climates 69 have been yet proposed. The lack of a general understanding on how benthic macroinvertebrate 70 communities respond across the hydrological gradient in floodplains may be partly due to a lack of 71 comparable field observations concerning their distribution from a range of rivers covering a broad 72 gradient of climatic and hydrological conditions (such as in Minshall et al. 1983). 73 In an attempt to provide a framework for comparing ecological processes and responses between 74 different river systems, a number of authors have undertaken river classifications from different regions 75 (e.g. Kennard et al. 2010, Poff and Ward 1989). Broadly, these classifications have identified groups of 76 rivers that range from perennial rivers and streams, characterised by consistent base flow and 77 predictable flood patterns –and often represented in temperate regions—to intermittent rivers and 78 streams that may have variable periods of cease to flow but can flood predictably. This latter case 79 concerns many Mediterranean and subtropical systems (e.g. Bonada et al. 2007, Depetris 2007), 80 whereas unpredictable floods occur in many arid and semi-arid systems (e.g.Kennard et al. 2010). Such 3 81 differences in climate and hydrology should result in different spatio-temporal floodplain conditions 82 yielding differences in taxonomic structure of aquatic communities (e.g. Poff and Ward 1990, Puckridge 83 et al. 1998). Owing to different taxonomic representations across broad regional scales, it can be 84 difficult to get a clear picture of similarities or general patterns if just relying on taxonomic comparisons. 85 In contrast, species traits across broad regions may provide insight into general patterns related to 86 hydrological connectivity, since traits aggregate biological information shared among taxonomically 87 different taxa. Therefore, while large differences in taxonomic composition are expected for rivers from 88 distinct climate or biogeographic regions, similarity in trait response across the hydrological gradient is 89 anticipated (e.g. Charvet et al. 2000). 90 Although many authors have reported a significant response of benthic macroinvertebrates to a 91 gradient of lateral connectivity, most of the available literature focuses on temperate European rivers, 92 such as the Rhône (e.g. Amoros and Bornette 2002, Castella et al. 1984, Paillex et al. 2007), the Danube 93 (e.g. Reckendorfer et al. 2006, Tockner et al. 1999, Ward et al. 1999) and the Rhine (e.g. Ward et al. 94 1999). Fewer studies exist in Mediterranean (but see Bonada et al. 2006, Gallardo et al. 2008), arid (but 95 see Sheldon et al. 2002, Sheldon and Thoms 2006), or subtropical rivers (but see Arrington and 96 Winemiller 2006, Marchese and Ezcurra de Drago 1992, Zilli et al. 2008). According to these and other 97 publications, EPT richness (Ephemeroptera, Plecoptera and Trichoptera) usually peaks in habitats with 98 high hydrological connectivity because of the high dissolved oxygen requirement and perhaps lower 99 thermal tolerance of associated species (Usseglio-Polatera and Tachet 1994). In contrast, COH richness 100 (Coleoptera, Odonata and Heteroptera) tends to peak in sites most disconnected from main channels 101 (e.g. Arscott et al. 2005, Bonada et al. 2006, Skern et al. 2010). Species richness and diversity usually 102 peak under intermediate hydrological disturbance conditions in accordance with the Intermediate 103 Disturbance Hypothesis (Connell 1978). Because of intense abiotic constraints, we might expect low trait 104 richness and diversity under high hydrological connectivity, while stability in intermediately connected 105 and fragmented channels may foster biotic interaction, thus trait richness (Statzner et al. 2004). 106 According to the River Habitat Templet (RHT, Townsend and Hildrew 1994), we might expect higher 107 proportions of small-sized individuals, associated with plurivoltinism and short life spans in the main 108 river channel submitted to frequent flooding. It is worth noting that highly variable flow regimes and 4 109 prolonged periods of extended hydrological
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